Human motor-unit recruitment during isometric contractions and repeated dynamic movements

Spike-triggered averaging was used to determine the twitch tensions and contraction times of motor units in the abductor pollicis brevis muscle of two human subjects for two directions of isometric contraction: abduction and opposition of the thumb. During isometric contractions in each direction, the threshold force for motor-unit recruitment and the twitch amplitude were correlated linearly. These data suggested that an orderly pattern of recruitment, according to increasing twitch size, describes the function of the human abductor pollicis brevis muscle for each contraction direction. Rank order of motor-unit recruitment in each isometric contraction direction was correlated, but not identical. All units contributed tension in each direction of contraction, so no clear evidence was found for task-dependent motor units. In two subjects, motor-unit recruitment order during isometric contraction of the first dorsal interosseous and abductor pollicis brevis muscles was then compared with that of motor-unit pairs in both muscles during repetitive dynamic movements. Recruitment according to increasing twitch size was largely preserved during the repetitive opening and closing of scissors. The recruitment reversals that were observed were usually between pairs of units with similar thresholds.     

The thenar muscles

Examination of the thenar muscles in 30 anatomical preparations of the hand have shown that the abductor pollicis brevis, the opponens pollicis, and the adductor pollicis muscles are made up of several muscle bellies. The number and insertions of these bellies are varied. Both heads of flexor pollicis brevis do not originate from any particular muscle belly. The superficial head of this muscle always inserted into the head of the thumb metacarpal, either completely, or, some of the fibres of the dorsal aponeurosis of the thumb were attached to the base of the proximal phalanx. Furthermore the anatomy of the abductor pollicis brevis muscle was related to the presence of a tendinous slip from abductor pollicis longus. These variations could have an influence on proprioception in the thumb ray.     

Characteristics of synergic relations during isometric contractions of human elbow muscles

We studied the patterns of EMG activity in elbow muscles in three normal human subjects. The myoelectrical activity of 7-10 muscles that act across the human elbow joint was simultaneously recorded with intramuscular electrodes during isometric joint torques exerted over a range of directions. These directions included flexion, extension, varus (internal humeral rotation), valgus (external humeral rotation), and several intermediate directions. The forces developed at the wrist covered a range of 360 degrees, all orthogonal to the long axis of the forearm. The levels of EMG activity were observed to increase with increasing joint torque in an approximately linear manner. All muscles were active for ranges less than 360 degrees and most were active for less than 180 degrees. The EMG activity was observed to vary in a systematic manner with changes in torque direction and, when examined over the full angular range at a variety of torque levels, is simply scaled with increasing torque magnitude. There were no torque directions or torque magnitudes for which a single muscle was observed to be active alone. In all cases, joint torque appeared to be produced by a combination of muscles. The direction for which the EMG of a muscle reached a maximum value was observed to correspond to the direction of greatest mechanical advantage as predicted by a simple mechanical model of the elbow and relevant muscles. Muscles were relatively inactive during varus torques. This implies that the muscles were not acting to stabilize the joint in this direction and could have been allowing ligaments to carry the load. Plots of EMG activity in one muscle against EMG activity in another demonstrate some instances of pure synergies, but patterns of coactivation for most muscles are more complicated and vary with torque direction. The complexity of these patterns raises the possibility that synergies are determined by the task and may have no independent existence. Activity in two heads of triceps brachii (medial head--a single-joint muscle and long head--a two-joint muscle) covaried closely for a range of torque magnitudes and directions, though shoulder torque and hence the forces experienced by the long head of the triceps undoubtedly varied. The similarity of activation patterns indicates that elbow torque was the principal determining factor. The origins of muscle synergies are discussed. It is suggested that they are best understood on the basis of a model which encodes limb torque in premotor neurons.(ABSTRACT TRUNCATED AT 400 WORDS)     

A dynamic model of the hand with application in functional neuromuscular stimulation

Potential hand function in people with tetraplegia was evaluated using a three-dimensional dynamic mathematical model. The model was used to evaluate hand positioning, grasp force, and the outcome of surgeries such as tendon transfers and joint fusion, in situations typical of those encountered when using functional neuromuscular stimulation to restore function in people with tetraplegia. In the model, the hand is treated as a jointed multibody system. Each joint is subjected to muscle moments, passive joint moment, and moments due to grasp forces. Model simulations showed that function was highly dependent on both muscle strength and joint passive moments. The potential for tendon transfers, such as the Zancolli-lasso and intrinsic-plasty, to improve hand function was demonstrated, but their value is subject-dependent. It was also shown that activation of multiple thumb muscles (adductor pollicis, abductor pollicis brevis, and flexor pollicis longus) without interphalangeal joint fusion can provide convenient lateral pinch posture with ∼70% more grip force than a currently used method, which includes joint fusion but requires only one muscle. Finally, a grasp protocol was introduced and shown successful in palmar grasp and hold of movable cylindrical objects using only extrinsic muscles, provided the fingers could be extended sufficiently to enclose the object.     

An Analysis of Extraocular Muscle Forces in the Piked Dogfish (Squalus acanthias)

Vertebrates utilize six extraocular muscles that attach to a tough, protective sclera to rotate the eye. The goal of the study was to describe the maximum tetanic forces, as well as the torques produced by the six extraocular muscles of the piked dogfish Squalus acanthias to understand the forces exerted on the eye. The lateral rectus extraocular muscle of Squalus acanthias was determined to be parallel fibered with the muscle fibers bundled into discrete fascicles. The extraocular muscles attach to the sclera by muscular insertions. The total tensile forces generated by the extraocular muscles ranged from 1.18 N to 2.21 N. The torques of the extraocular muscles ranged from 0.39 N to 2.34 N. The torques were greatest in the principal direction of movement for each specific muscle. The lateral rectus produced the greatest total tensile force, as well as the greatest torque force component, while the medial rectus produced the second greatest. This is likely due to the constant rotational movement of the eye anteriorly and posteriorly to stabilize the visual image, as well as increase the effective visual field during swimming. Rotational forces in dimensions other than the primary direction of movement may contribute to motion in directions other than the principal direction during multi-muscle contraction that occurs in the vertebrate eye. Anat Rec, 2018. © 2018 Wiley Periodicals, Inc. Anat Rec, 302:837–844, 2019. © 2018 Wiley Periodicals, Inc.     

Common input to motor units of digit flexors during multi-digit grasping

The control of whole hand grasping relies on complex coordination of multiple forces. While many studies have characterized the coordination of finger forces and torques, the control of hand muscle activity underlying multi-digit grasping has not been studied to the same extent. Motor-unit synchrony across finger muscles or muscle compartments might be one of the factors underlying the limited individuation of finger forces. Such "unwanted" coupling among finger forces, however, might be desirable when a high level of force coupling is required to prevent object slip during grasping. The goal of this study was to quantify the strength of synchrony between single motor units from extrinsic hand muscles as subjects held a device with a five-digit grasp. During the hold phase, we recorded the normal force exerted by each digit and the electrical activity of single motor units from each of the four divisions of the muscle flexor digitorum profundus (FDP) and one thumb flexor muscle, m. flexor pollicis longus (FPL). The strength of motor-unit synchrony was quantified by the common input strength index (CIS). We found moderate to strong motor-unit synchrony between FPL and the index FDP compartment [CIS: 0.49 +/- 0.03 (SE)] and across most FDP compartments (0.34 +/- 0.02). Weak synchrony, however, was found between FPL and the middle, ring, and little finger FDP compartments (0.25 +/- 0.01). This difference might reflect the larger force contribution of the thumb-index finger pair relative to other thumb-finger combinations in five-digit grasping.     

人鱼际肌的肌内神经分支分布和肌梭密度研究

目的探索鱼际肌肌肌内神经分支和肌梭密度的分布。方法采用改良Sihler’s肌内神经染色法和HE染色法进行解剖学研究。结果鱼际肌的神经常从肌起端深面入肌,神经入肌后在拇短展肌、拇对掌肌、拇收肌横头内与肌长轴垂直走向,拇收肌斜头和拇短屈肌内沿肌长轴平行走形。80%～82.5%的拇短屈肌和拇指对掌肌接受正中神经和尺神经的双重支配。拇短屈肌浅头和深头、拇收肌横头和斜头有独立的神经支配,可分出神经肌肉亚部。4块肌内神经分支分布密集区多在肌的中部与近端,可见"Y"、"O"、"H"或"U"型等不同的神经吻合形式。鱼际肌肌梭密度高达16.19～27.14个/g,高低顺序为拇指对掌肌拇短屈肌拇短展肌拇收肌。结论鱼际肌肌内神经吻合丰富,肌梭密度高,除拇对掌肌外,其余肌块可作整肌或半肌移植的供体。     

Characteristics of the electrophysiological activity of muscles attached to the transverse carpal ligament in carpal tunnel syndrome

The main cause of carpal tunnel syndrome (CTS) remains unknown. Stiffness of the subcutaneous area of the volar aspect of the carpal tunnel is present in many patients and suggests that the stiffness of muscles attached to the transverse carpal ligament is increased. We performed an electrophysiological study to investigate muscle activities and to clarify whether the stiffness of muscles attached to the transverse carpal ligament is involved in the pathogenesis of CTS. The subjects of this study included 16 patients with early CTS showing no motor dysfunction. Both thenar muscles (opponens pollicis, abductor pollicis brevis, and flexor pollicis brevis) and hypothenar muscles (opponens digiti minimi, abductor digiti minimi, flexor digiti minimi brevis) were investigated. Surface electrodes were placed on each muscle, and maximum voluntary contractions with the thumb and little finger in opposition were maintained for 3 seconds in all patients and in 7 control subjects. Electromyographs were subjected to fast Fourier transform analysis, and the root mean square (RMS) and the mean power frequency (MPF) were determined for each muscle. The RMS of the opponens pollicis was significantly less in hands affected by CTS (292.8 μV) than in healthy hands (405.9 μV). The RMS did not differ between affected hands and healthy hands for the other 2 thenar muscles but did differ significantly for the hypothenar muscles. The MPF did not differ between affected hands and healthy hands for any muscle. The results show that electrophysiological differences are present among muscles innervated by the median nerve and that hypothenar muscles originally unrelated to median nerve dysfunction are also affected in early CTS. These results suggest that modulation of muscles attached to the transverse carpal ligament is involved in the pathogenesis of CTS.     

Diminished capacity to modulate motor activation patterns according to task contributes to thumb deficits following stroke

The objective of this study was to explore motor impairment of the thumb following stroke. More specifically, we quantitatively examined kinetic deficits of the thumb. We anticipated that force deficits would be nonuniformly distributed across the kinetic workspace, due in part to varying levels of difficulty in altering the motor activation pattern to meet the task. Eighteen stroke survivors with chronic hemiparesis participated in the trials, along with nine age-matched controls. Of the stroke-survivor group, nine subjects had moderate hand impairment, and the other nine subjects had severe hand impairment. Subjects were instructed to generate maximal isometric thumb-tip force, as measured with a load cell, in each of six orthogonal directions with respect to the thumb tip. Activity of three representative thumb muscles was monitored through intramuscular and surface electrodes. Univariate split-plot analysis of variance revealed that clinical impairment level had a significant effect on measured force (P < 0.001), with the severely impaired group producing only 13% of the control forces, and the moderately impaired group generating 32% of control forces, on average. Weakness in the moderately impaired group exhibited a dependence on force direction (P = 0.015), with the least-relative weakness in the medial direction. Electromyographic recordings revealed that stroke survivors exhibited limited modulation of thumb-muscle activity with intended force direction. The difference in activation presented by the control group for a given muscle was equal to 40% of its full activation range across force directions, whereas this difference was only 26% for the moderately impaired group and 15% for the severely impaired group. This diminished ability to modify voluntary activation patterns, which we observed previously in index-finger muscles as well, appears to be a primary factor in hand impairment following stroke.     

拇短展肌对拇指指骨间关节背伸作用的解剖学研究

目的　为解决胸小肌移植重建拇对掌功能术后出现拇指指骨间关节屈曲畸形提供解剖学依据并寻找解决办法。 方法　在15具尸体手标本上,使拇指处于对掌位时,测量拇指指骨间关节在拇短展肌未切断及切断两种情况下的自然屈曲角度;同时,通过外力作用使拇指指骨间关节分别屈曲处于45°及60°时,分析拇短展肌对拇指指骨间关节屈伸功能的影响。 结果　拇指处于对掌位时,拇短展肌在未切断及切断两种情况下,拇指指骨间关节自然屈曲角度分别为（13.30±2.13）°及（24.03±1.25）°;同时,使拇指指骨间关节屈曲处于45°时,所用外力分别约（0.50±0.08）N及（0.22±0.07）N,而使拇指指骨间关节屈曲处于60°时,所用外力分别约（1.48±0.09）N及（1.15±0.04）N。 结论　拇短展肌对拇指指骨间关节主要起背伸作用,因此胸小肌移植重建拇对掌功能时,应同时重建拇短展肌功能,才有可能纠正在拇对掌功能重建术后出现的拇指指骨间关节屈曲畸形。     

Progressive suppression of intracortical inhibition during graded isometric contraction of a hand muscle is not influenced by hand preference

GABAergic intracortical inhibition (ICI) in human motor cortex (M1) assists fractionated activation of muscles, and it has been suggested that hemispheric differences in ICI may contribute to hand preference. Previous studies of this issue have all been conducted at rest, with conflicting results. Testing during voluntary activation may reveal functionally relevant differences. In normal subjects, we assessed (1) operation of ICI circuits during selective activation of an intrinsic hand muscle at different forces, and (2) whether this differs between right and left hemispheres. Surface EMG was recorded bilaterally from abductor pollicis brevis (APB), first dorsal interosseous (FDI) and abductor digiti minimi (ADM) muscles in eleven right-handed subjects. A circular coil applied paired transcranial magnetic stimulation (TMS) with posteriorly directed current in the brain. Conditioning intensity was 0.8 × active threshold and interstimulus interval was 3 ms. TMS was applied to right or left M1 while subjects were at rest or performing isometric thumb abduction at different forces (0.5, 1, 2, 3, 5 and 10 N) with the contralateral hand. Conditioning TMS was less effective at suppressing the muscle evoked potential in APB during 2–10 N thumb abduction (P < 0.0001) versus rest, but not with lower target forces (0.5, 1 N). Conditioning TMS was less effective for FDI and ADM only during 10 N thumb abduction. We conclude that differential modulation of ICI in M1 during selective muscle activation is a function of target isometric force level. At low forces (<5% MVC), ICI is not modulated for the corticospinal neurons controlling the active or inactive muscles. There is a progressive reduction of ICI effects on corticospinal neurons at higher forces, which is largely restricted to corticospinal neurons controlling the muscle targeted for activation over the range of forces tested (up to ∼25% MVC). The pattern of ICI modulation with selective voluntary muscle contraction was similar in left and right hemispheres during this relatively simple static task. If hemispheric differences in operation of M1 ICI circuits contribute to hand preference, a more challenging finger movement protocol may be needed to demonstrate this asymmetry.     

Biomechanical alterations in the carpal arch and hand muscles after carpal tunnel release: A further approach toward understanding the function of the flexor retinaculum and the cause of postoperative grip weakness

The difference between maximal and minimal distance covered (the distance between the trapezium ridge and hamate hook; moment exerted on structures: I Nm) by an intact flexor retinaculum (FR; minimum, 3.3 ± 0.1 cm; maximum, 3.7 ± 0.2 cm) and the increase in the maximal distance on carpal tunnel release (CTR; increase, 1.6 ± 0.2 mm) were significant. Under an external supination moment, the distance between the attachments of the trapeziopisiform band increased after CTR. Under external pronation and ulnar abduction moments, the distance between the attachments of the scaphoideohamate band increased after CTR. The CTR resulted in an anatomic attachment loss for the following muscles: the superficial head of the flexor pollicis brevis (shortening by ∼25%, relative to rest length), the ulnar part of the abductor pollicis brevis (with opposition and adductory functions, ∼20%), the opponens pollicis (∼20%), the middle part of the abductor pollicis brevis (∼7%), and the opponens digiti minimi (∼10%). Preoperative and postoperative (2–7 weeks after surgery) measurements of the reaction force of the distal phalanx (under isometric thumb opposition and finger II–IV flexion with extended carpal joint) led to differentiation of three groups: (1) significant strength loss—the patients showed difficulties with grasping, lifting, twisting off lids and caps, screwing, pulling ropes, and pinching; (2) no significant change in force values; and (3) a significant increase in strength (patients who could grip more firmly). © 1996 Wiley-Liss, Inc.     

Time course of determination of movement direction in the reaction time task in humans

The primary motor cortex produces motor commands that include encoding the direction of movement. Excitability of the motor cortex in the reaction time (RT) task can be assessed using transcranial magnetic stimulation (TMS). To elucidate the timing of the increase in cortical excitability and of the determination of movement direction before movement onset, we asked six right-handed, healthy subjects to either abduct or extend their right thumb after a go-signal indicated the appropriate direction. Between the go-signal and movement onset, single TMS pulses were delivered to the contralateral motor cortex. We recorded the direction of the TMS-induced thumb movement and the amplitude of motor-evoked potentials (MEPs) from the abductor pollicis brevis and extensor pollicis brevis muscles. Facilitation of MEPs from the prime mover, as early as 200 ms before the end of the reaction time, preceded facilitation of MEPs from the nonprime mover, and both preceded measurable directional change. Compared with a control condition in which no voluntary movement was required, the direction of the TMS-induced thumb movement started to change in the direction of the intended movement as early as 90 ms before the end of the RT, and maximum changes were seen shortly before the end of reaction time. Movement acceleration also increased with maxima shortly before the end of the RT. We conclude that in concentric movements a change of the movement direction encoded in the primary motor cortex occurs in the 200 ms prior to movement onset, which is as early as increased excitability itself can be detected.     

Control of grip force during restraint of an object held between finger and thumb: responses of muscle and joint afferents from the digits

Pulling or pushing forces applied to an object gripped between finger and thumb excite tactile afferents in the digits in a manner awarding these afferents probable roles in triggering the reactive increases in grip force and in scaling the changes in grip force to the changes in applied load-force. In the present study we assessed the possible contributions from slowly adapting afferents supplying muscles involved in the generation of grip forces and from digital joint afferents. Impulses were recorded from single afferents via tungsten microelectrodes inserted percutaneously into the median or ulnar nerves of awake human subjects. The subject held a manipulandum with a precision grip between the receptor-related digit (index finger, middle finger, ring finger or thumb) and an opposing digit (thumb or index finger). Ramp-and-hold load forces of various amplitudes (0.5–2.0 N) and ramp rates (2–32 N/s) were delivered tangential to the parallel grip surfaces in both the distal (pulling) and the proximal (pushing) directions. Afferents from the long flexors of the digits (n=19), regardless of their muscle-spindle or tendon-organ origin, did not respond to the load forces before the onset of the automatic grip response, even with the fastest ramp rates. Their peak discharge closely followed the peak rate of increase in grip force. During the hold phase of the load stimulus, the afferents sustained a tonic discharge. The discharge rates were significantly lower with proximally directed loads despite the mean grip-force being similar in the two directions. This disparity could be explained by the differing contributions of these muscles to the finger-tip forces necessary to restrain the manipulandum in the two directions. Most afferents from the short flexors of the digits (n=17), including the lumbricals, dorsal interossei, opponens pollicis, and flexor pollicis brevis, did not respond at all, even with the fastest ramps. Furthermore, the ensemble pattern from the joint afferents (n=6) revealed no significant encoding of changes in finger-tip forces before the onset of the increase in grip force. We conclude that mechanoreceptors in the flexors of the digits and in the interphalangeal joints cannot be awarded a significant role in triggering the automatic changes in grip force. Rather, their responses appeared to reflect the reactive forces generated by the muscles to restrain the object. Hence, it appears that tactile afferents of the skin in contact with the object are the only species of receptor in the hand capable of triggering and initially scaling an appropriate change in grip force in response to an imposed change in load force, but that muscle and joint afferents may provide information related to the reactive forces produced by the subject.     

Force path curvature and conserved features of muscle activation

This study examined the patterns of muscle activity that subserve the production of dynamic isometric forces in various directions. The isometric condition provided a test for basic features of neuromuscular control, since the task was analogous to reaching movement, but the behavior was not necessarily shaped by the anisotropy of inertial and viscoelastic resistance to movement. Electromyographic (EMG) activity was simultaneously recorded from nine elbow and/or shoulder muscles, and force pulses, steps, and ramps were monitored using a transducer fixed to the constrained wrists of human subjects. The force responses were produced by activating shoulder and elbow muscles; response direction was controlled by the relative intensity of activity in muscles with different mechanical actions. The primary objective was to characterize the EMG temporal pattern. Ideally, synchronous patterns of phasic muscle activation (and synchronous dynamic elbow and shoulder torques) would result in a straight force path; asynchronous muscle activation could result in substantial force path curvature. For both pulses and steps, asynchronous muscle activation was observed and was accompanied by substantial force path curvature. A second objective was to compare phasic and tonic EMG activity. The spatial tuning of EMG intensity was similar for the phasic and tonic activities of each muscle and also similar to the spatial tuning of tonic activity in a previous study where the arm was stationary but unconstrained.     

Strategies that simplify the control of quadrupedal stance. II. Electromyographic activity

1. This study tested the hypothesis that muscle synergies underlie the invariance in the direction of corrective forces observed following stance perturbations in the horizontal plane. Electromyographic activity was recorded from selected forelimb and hindlimb muscles of cats subjected to horizontal translations of the supporting surface in 16 different directions. The responses of muscles were quantified for each perturbation, and tuning curves were constructed that related the amplitude of muscle response to the direction of platform movement. 2. Muscle tuning curves tended to group into one of two regions, corresponding to the two directions of force vectors. A few muscles showed clearly different recruitment patterns. The same direction of correction force vector was produced by different patterns of muscle activity, and the particular EMG pattern depended on the direction of platform movement. Therefore a simple muscle synergy organization could not account for the invariance in force vector generation. 3. It is concluded that there is a hierarchy of control in the maintenance of stance in which the vector of force exerted against the ground is a high level, task-dependent controlled variable and the selection of muscles to activate in order to produce the vector is controlled at a lower level. It is proposed that muscles are controlled using a modified synergy strategy. In this scheme, a synergy is not simply a fixed group of muscles, constrained to act as a unit. Rather, muscles are organized as a task-dependent synergy that is tuned or modified as needed by the addition or subtraction of other muscles.     

Biometry of thenar muscle origins on the flexor retinaculum

The transverse carpal ligament (TCL), the main part of the flexor retinaculum, serves as an anchor for the thenar muscles: abductor pollicis brevis (APB), superficial head of the flexor pollicis brevis (sFPB), and opponens pollicis (OPP). Biomechanically, the thenar muscles rely on their TCL anchoring to transmit muscle contractions distally for thumb force and motion production, and reciprocally, muscle contraction interacts with the TCL at the proximal end through the origins. However, scarce knowledge exists regarding the distribution pattern of the thenar muscle origins. The purpose of this study was to understand the anatomical interface between the thenar muscles and TCL by examining the origin distributions of the individual muscles. Ten cadaveric specimens were dissected for digitization of the muscle origins and TCL volar surface. Digitized data were used for mesh reconstruction and calculation of surface areas and centroids. The origin areas for APB, sFPB, and OPP were 105.8 ± 30.3, 64.6 ± 15.2, and 245.9 ± 70.7 mm², respectively. The surface area of the TCL was 386.2 ± 86.9 mm². The origin areas of APB and OPP on the TCL were comparable, 18.4 ± 4.8% and 17.3 ± 9.6% of the TCL area, respectively. The origin locations for APB, sFPB, and OPP were in proximal-radial quadrant of the TCL, on distal aponeurosis outside the TCL, and around the ridge of trapezium, respectively. The knowledge of the anatomical interface provides a foundation for the understanding of biomechanical interactions between the muscles and ligaments and pathomechanical implications.     

Control of the wrist in three-joint arm movements to multiple directions in the horizontal plane

In a reaching movement, the wrist joint is subject to inertial effects from proximal joint motion. However, precise control of the wrist is important for reaching accuracy. Studies of three-joint arm movements report that the wrist joint moves little during point-to-point reaches, but muscle activities and kinetics have not yet been described across a range of movement directions. We hypothesized that to minimize wrist motion, muscle torques at the wrist must perfectly counteract inertial effects arising from proximal joint motion. Subjects were given no instructions regarding joint movement and were observed to keep the wrist nearly motionless during center-out reaches to directions throughout the horizontal plane. Consistent with this, wrist muscle torques exactly mirrored interaction torques, in contrast to muscle torques at proximal joints. These findings suggest that in this reaching task the nervous system chooses to minimize wrist motion by anticipating dynamic inertial effects. The wrist muscle torques were associated with a direction-dependent choice of muscles, also characterized by initial reciprocal activation rather than initial coactivation to stiffen the wrist joint. In a second experiment, the same pattern of muscle activities persisted even after many trials reaching with the wrist joint immobilized. These results, combined with similar features at the three joints, such as cosine-like tuning of muscle torques and of muscle onsets across direction, suggest that the nervous system uses similar rules for muscles at each joint, as part of one plan for the arm during a point-to-point reach.     

The innervation of deep muscles of the human forearm extensors]

The innervation of four deep muscles of the human forearm extensors (the abductor pollicis longus, the extensor pollicis brevis, the extensor pollicis longus, and the extensor indicis muscles) were investigated in 24 bodies (48 sides) from those used in the 1989 and 1990 student courses in gross anatomy dissection at the Iwate Medical University School of Medicine. The forearm extensor muscles and the deep branch of the radial nerve were dissected intensively in the student courses in gross anatomy and were removed afterwards. The four deep muscles of the human forearm extensors and the nerves innervating the muscles were observed while they were immersed in the water and with use of a stereomicroscope--with the assistance of which they were drawn. In six sides the intramuscular nerve supply was also examined carefully and drawn. The results were as follows. 1. The nerves to the four deep muscles of the forearm extensors arose usually from the deep branch of the radial nerve after emerging the supinator muscle and sending branches to superficial forearm extensors. In some cases a nerve or nerves to the superficial forearm extensors were observed arising from the deep branch of the radial nerve after sending one or more branches to the deep forearm extensor muscles, or from the branches to the deep muscles themselves. However they were split easily from the deep branch of the radial nerve and from the branches to the four deep forearm extensors proximally near to the emerging of the deep branch from the supinator muscle. Therefore, it was considered to be constant that the nerves to the four deep forearm extensors arose from the deep branch of the radial nerve after branching to the superficial forearm extensors. 2. The radial group of the deep forearm extensors (the abductor pollicis longus and the extensor pollicis brevis muscles) was innervated usually by one branch that arose from the deep branch of the radial nerve just after emerging from the supinator and giving off branches to the superficial forearm extensors. This branch ran on the dorsal (extensor) surface of the abductor pollicis longus muscle distally, sending many twigs to this muscle, and entered into the muscle at various distances from the origin (Figs. 1-6). The abductor pollicis brevis muscle was innervated by some twigs that ran usually inside but occasionally outside of the abductor pollicis longus muscle (Figs. 7-10).(ABSTRACT TRUNCATED AT 400 WORDS)